Dynamics and stimulus-dependence of pacemaker control during behavioral modulations in the weakly electric fish,Apteronotus

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Abstract
Weakly electric fish generate around their bodies low-amplitude, AC electric fields which are used both for the detection of objects and intraspecific communication. The types of modulation in this signal of which the high-frequency wave-type gymnotiform,Apteronotus, is capable are relatively few and stereotyped. Chief among these is the chirp, a signal used in courtship and agonistic displays. Chirps are brief and rapid accelerations in the normally highly regular electric organ discharge (EOD) frequency. Chirping can be elicited artificially in these animals by the use of a stimulus regime identical to that typically used to elicit another behavior, the jamming avoidance response (JAR). The neuronal basis for the JAR, a much slower and lesser alteration in EOD frequency, is well understood. Examination of the stimulus features which induce chirping show that, like the JAR, there is a region of frequency differences between the fish's EOD and the interfering signal that maximally elicits the response. Moreover, the response is sex-specific with regard to the sign of the frequency difference, with females chirping preferentially on the positive and most males on the negative Df. These features imply that the sensory mechanisms involved in the triggering of these communicatory behaviors are fundamentally similar to those explicated for the JAR. Additionally, two other modulatory behaviors of unknown significance are described. The first is a non-selective rise in EOD frequency associated with a JAR stimulus, occurring regardless of the sign of the Df. This modulation shares many characteristics with the JAR. The second behavior, which we have termed a ‘yodel’, is distinct from and kinetically intermediate to chirping and the JAR. Moreover, unlike the other studied electromotor behaviors it is generally produced only after the termination of the eliciting stimulus.